Physical activity and temperature changes of Asian elephants (Elephas maximus) participating in eco-tourism activities and elephant polo.

Captive and domestic animals are often required to engage in physical activity initiated or organised by humans, which may impact their body temperature, with consequences for their health and welfare. This is a particular concern for animals such as elephants that face thermoregulatory challenges because of their body size and physiology. Using infrared thermography, we measured changes in skin temperature associated with two types of physical activity in ten female Asian elephants (Elephas maximus) at an eco-tourism lodge in Nepal. Six elephants took part in an activity relatively unfamiliar to the elephants-a polo tournament-and four participated in more familiar ecotourism activities. We recorded skin temperatures for four body regions affected by the activities, as well as an average skin temperature. Temperature change was used as the response variable in the analysis and calculated as the difference in elephant temperature before and after activity. We found no significant differences in temperature change between the elephants in the polo-playing group and those from the non-polo playing group. However, for both groups, when comparing the average skin body temperature and several different body regions, we found significant differences in skin temperature change before and after activity. The ear pinna was the most impacted region and was significantly different to all other body regions. This result highlights the importance of this region in thermoregulation for elephants during physical activity. However, as we found no differences between the average body temperatures of the polo and non-polo playing groups, we suggest that thermoregulatory mechanisms can counteract the effects of both physical activities the elephants engaged in.

Multimodal communication and audience directedness in the greeting behaviour of semi-captive African savannah elephants.

Many species communicate by combining signals into multimodal combinations. Elephants live in multi-level societies where individuals regularly separate and reunite. Upon reunion, elephants often engage in elaborate greeting rituals, where they use vocalisations and body acts produced with different body parts and of various sensory modalities (e.g., audible, tactile). However, whether these body acts represent communicative gestures and whether elephants combine vocalisations and gestures during greeting is still unknown. Here we use separation-reunion events to explore the greeting behaviour of semi-captive elephants (Loxodonta africana). We investigate whether elephants use silent-visual, audible, and tactile gestures directing them at their audience based on their state of visual attention and how they combine these gestures with vocalisations during greeting. We show that elephants select gesture modality appropriately according to their audience's visual attention, suggesting evidence of first-order intentional communicative use. We further show that elephants integrate vocalisations and gestures into different combinations and orders. The most frequent combination consists of rumble vocalisations with ear-flapping gestures, used most often between females. By showing that a species evolutionarily distant to our own primate lineage shows sensitivity to their audience's visual attention in their gesturing and combines gestures with vocalisations, our study advances our understanding of the emergence of first-order intentionality and multimodal communication across taxa.

Maximum trunk tip force assessment related to trunk position and prehensile 'fingers' implication in African savannah elephants.

African elephants have a wide range of abilities using their trunk. As a muscular hydrostat, and thanks to the two finger-like processes at its tip, this proboscis can both precisely grasp and exert considerable force by wrapping. Yet few studies have attempted to quantify its distal grasping force. Thus, using a device equipped with force sensors and an automatic reward system, the trunk tip pinch force has been quantified in five captive female African savanna elephants. Results showed that the maximum pinch force of the trunk was 86.4 N, which may suggest that this part of the trunk is mainly dedicated to precision grasping. We also highlighted for the first time a difference in force between the two fingers of the trunk, with the dorsal finger predominantly stronger than the ventral finger. Finally, we showed that the position of the trunk, particularly the torsion, influences its force and distribution between the two trunk fingers. All these results are discussed in the light of the trunk's anatomy, and open up new avenues for evolutionary reflection and soft robot grippers.

Characteristics of gut microbiota in captive Asian elephants (Elephas maximus) from infant to elderly.

Gut microbiota play an important role in the health and disease of Asian elephants, however, its characteristics at each stage of life have not been thoroughly investigated in maintaining and regulating health of elephants. This study, therefore, aimed to characterize the profiles of the gut microbiota of captive Asian elephants from infants to the elderly. Gut microbiota were identified by 16S rRNA sequencing from the feces of captive Asian elephants with varying age groups, including infant calves, suckling calves, weaned calves, subadult and adult elephants, and geriatric elephants. The diversity of the gut microbiota was lowest in infants, stable during adulthood, and slightly decreased in the geriatric period. The gut microbiota of the infant elephants was dominated by milk-fermenting taxa including genus Bifidobacterium of family Bifidobacteriaceae together with genus Akkermansia. The fiber-fermenting taxa such as Lachnospiraceae_NK3A20_group were found to be increased in suckling elephants in differential abundance analysis by Analysis of Compositions of Microbiomes with Bias Correction (ANCOM-BC). The gut microbiota profiles after weaning until the adult period has been uniform as indicated by no significant differences in beta diversity between groups. However, the composition of the gut microbiota was found to change again in geriatric elephants. Understanding of the composition of the gut microbiota of captive Asian elephants at various life stages could be beneficial for promoting good health throughout their lifespan, as well as ensuring the welfare of captive elephants.

Asian elephants distinguish sexual status and identity of unfamiliar elephants using urinary odours.

Despite the ubiquity of odours in mammals, few studies have documented the natural olfactory abilities of many 'non-model' species such as the Asian elephant. As Asian elephants are endangered, we may apply odours to more effectively manage threatened populations. We implemented a habituation-discrimination paradigm for the first time in Asian elephants to test the ability of elephants to discriminate between unfamiliar male elephant urine, hypothesizing that elephants would successfully distinguish non-musth from musth urine and also distinguish identity between two closely related individuals. We conducted two bioassay series, exposing three female and three male zoo-housed elephants to the same urine sample (non-musth urine in the first series, and urine from an unfamiliar individual in the second) over 5 days. On the sixth day, we simultaneously presented each elephant with a novel sample (either musth urine or urine from a second unfamiliar individual) alongside the habituated urine sample, comparing rates of chemosensory response to each sample to indicate discrimination. All elephants successfully discriminated non-musth from musth urine, and also urine from two unfamiliar half-brothers. Our results further demonstrate the remarkable olfactory abilities of elephants with promising implications for conservation and management.

Identifying potential measures of stress and disturbance during a captive to wild African elephant reintegration.

There is increasing evidence of compromised welfare for elephants managed in captivity. Should such facilities eventually close, more elephants will need to be rehabilitated and reintegrated into the wild. The goal of such reintegration would be to restore any physical or psychological aspects of the elephant that may have been compromised in captivity, followed by introduction into a free-roaming system where they can interact with other elephants. However, to achieve this goal, the reintegration methods implemented need to be assessed to ensure that welfare remains the priority. The objective of this study was to test whether parameters generally associated with stress and disturbance in African elephants, respond to changes in potentially stressful environmental conditions, assessed at multiple temporal scales ranging from minutes to months. The main changes in environmental conditions that were investigated included the different phases of reintegration of a group of elephants from captivity into the wild. Stress and disturbance related parameters used for comparisons included physiological responses, namely the extent of temporal gland secretions (eTGS) and faecal glucocorticoid metabolite (fGCM) levels as well as behavioural responses, namely the display of stereotype and stress-related behaviours. Results showed that eTGS significantly increased during the initial release of the elephants compared to when in captivity. Stereotypic behaviours were only recorded during the captive phase and immediately ceased after release. Faecal GCM levels spiked in the first year after release before decreasing back to pre-reintegration levels during the third year. These findings indicate that fGCM levels, the eTGS and disturbance related behaviours all proved effective in explaining the changes in stress and disturbance experienced by elephants during the initial years after being reintegrated from captivity into the wild.

Frightened of giants: fear responses to elephants approach that of predators.

Animals are faced with a variety of dangers or threats, which are increasing in frequency with ongoing environmental change. While our understanding of fearfulness of such dangers is growing in the context of predation and parasitism risk, the extent to which non-trophic, interspecific dangers elicit fear in animals remains less appreciated. We provide an experimental test for fear responses of savannah ungulates to a dominant and aggressive megaherbivore, the African bush elephant (Loxodonta africana), and contrast responses to an apex predator known to elicit fear in this system. Using an automated behavioural response system, we contrast vigilance and run responses of ungulates to elephant, leopard (Panthera pardus), and control (red-chested cuckoo Cuculus solitarius) vocalizations. Overall, we find that ungulates responded to elephant calls, both in terms of an increase in run and vigilance responses relative to controls. The magnitude of most behavioural responses (four of six considered) to elephant vocalizations were not significantly different than responses to leopards. These results suggest that megaherbivores can elicit strong non-trophic fear responses by ungulates and call to broaden frameworks on fear to consider dominant species, such as megaherbivores, as key modifiers of fear-induced interactions.

Visual feedback influences the consistency of the locomotor pattern in Asian elephants (Elephas maximus).

Elephants are atypical of most quadrupeds in that they maintain the same lateral sequence footfall pattern across all locomotor speeds. It has been speculated that the preservation of the footfall patterns is necessary to maintain a statically stable support polygon. This should be a particularly important constraint in large, relatively slow animals. This suggests that elephants must rely on available sensory feedback mechanisms to actively control their massive pillar-like limbs for proper foot placement and sequencing. How the nervous system of elephants integrates the available sensory information for a stable gait is unknown. Here we explored the role that visual feedback plays in the control of the locomotor pattern in Asian elephants. Four Asian elephants (Elephas maximus) walked with and without a blindfold as we measured their stride time intervals. Coefficient of variation was used to assess changes in the overall variability of the stride time intervals, while approximate entropy was used to measure the stride-to-stride consistency of the time intervals. We show that visual feedback plays a role in the stride-to-stride consistency of the locomotor pattern in Asian elephants. These results suggest that elephants use visual feedback to correct and maintain proper sequencing of the limbs during locomotion.

Shape variation in the limb long bones of modern elephants reveals adaptations to body mass and habitat.

During evolution, several vertebrate lineages have shown trends towards an increase in mass. Such a trend is associated with physiological and musculoskeletal changes necessary to carry and move an increasingly heavy body. Due to their prominent role in the support and movement of the body, limb long bones are highly affected by these shifts in body mass. Elephants are the heaviest living terrestrial mammals, displaying unique features allowing them to withstand their massive weight, such as the columnarity of their limbs, and as such are crucial to understand the evolution towards high body mass in land mammals. In this study, we investigate the shape variation of the six limb long bones among the modern elephants, Elephas maximus and Loxodonta africana, to understand the effect of body mass and habitat on the external anatomy of the bones. To do so, we use three-dimensional geometric morphometrics (GMMs) and qualitative comparisons to describe the shape variation, at both the intraspecific and interspecific levels. Our results reveal that the two species share similar negative ontogenetic allometric patterns (i.e. becoming stouter with increased length) in their humerus and femur, but not in the other bones: the proximal epiphyses of the stylopod bones develop considerably during growth, while the distal epiphyses, which are involved in load distribution in the elbow and knee joints, are already massive in juveniles. We attribute this pattern to a weight-bearing adaptation already present in young specimens. Among adults of the same species, bone robustness increases with body mass, so that heavier specimens display stouter bones allowing for a better mechanical load distribution. While this robustness variation is significant for the humerus only, all the other bones appear to follow the same pattern. This is particularly visible in the ulna and tibia, but less so in the femur, which suggests that the forelimb and hindlimb adapted differently to high body mass support. Robustness analyses, while significant for the humerus only, suggest more robust long bones in Asian elephants than in African savanna elephants. More specifically, GMMs and qualitative comparisons indicate that three bones are clearly distinct when comparing the two species: in E. maximus the humerus, the ulna and the tibia display enlarged areas of muscular insertions for muscles involved in joint and limb stabilization, as well as in limb rotation. These results suggest a higher limb compliance in Asian elephants, associated with a higher dexterity, which could be linked to their habitat and foraging habits.

Elephant trunks use an adaptable prehensile grip.

Elephants have long been observed to grip objects with their trunk, but little is known about how they adjust their strategy for different weights. In this study, we challenge a female African elephant at Zoo Atlanta to lift 20-60 kg barbell weights with only its trunk. We measure the trunk's shape and wrinkle geometry from a frozen elephant trunk at the Smithsonian. We observe several strategies employed to accommodate heavier weights, including accelerating less, orienting the trunk vertically, and wrapping the barbell with a greater trunk length. Mathematical models show that increasing barbell weights are associated with constant trunk tensile force and an increasing barbell-wrapping surface area due to the trunk's wrinkles. Our findings may inspire the design of more adaptable soft robotic grippers that can improve grip using surface morphology such as wrinkles.

Predictors of testosterone in zoo-managed male African elephants (Loxodonta Africana).

Reproductive complications for both male and female zoo-managed African elephants (Loxodonta africana) contribute to the rapidly declining population. In zoo-managed bull elephants, few studies have explored the potential physiological, physical, social, and environmental factors that influence bull fertility, particularly, androgen production. Testosterone is the essential steroid hormone for male sexual maturation and inadequate concentrations can be detrimental for spermatogenesis. Testosterone, fecal glucocorticoid metabolites, leptin, glucose, insulin, and triglycerides were analyzed from weekly fecal and blood serum samples taken over 6 months from six zoo-managed African elephant bulls (10-19 years of age). Testosterone levels were compared to endocrine factors, weekly social and environmental variables, daily musth signs, and body condition scores (BCS). The glucose-to-insulin ratio (G:I) was the only physiological biomarker found to be positively associated with testosterone. Predictive physical variables included Musth Score (+) and Moderate Exercise (+). Bulls with BCS signifying overweight (BCS 4) had lower testosterone (36.6 ± 1.6 ng/g fecal extraction [FE]) than bulls with healthy BCS 3; 51.2 ± 4.9 ng/g FE). Numerous social variables influenced testosterone concentrations, including Total Contact Day (+), Female Interaction Day (+), Indirect Contact Day (+), Indirect Contact Night (+) and Total No Contact (-). Both percentage of Time Outdoor and Time Mixed positively influenced testosterone, whereas testosterone decreased for percentage of Time Indoors. Each additional daily browse opportunity increased testosterone by approximately 7 ng/g FE. In managed care, the emphasis should be placed on optimizing these predictors of testosterone production to promote bull reproductive health.

Seasonal range fidelity of a megaherbivore in response to environmental change.

For large herbivores living in highly dynamic environments, maintaining range fidelity has the potential to facilitate the exploitation of predictable resources while minimising energy expenditure. We evaluate this expectation by examining how the seasonal range fidelity of African elephants (Loxodonta africana) in the Kruger National Park, South Africa is affected by spatiotemporal variation in environmental conditions (vegetation quality, temperature, rainfall, and fire). Eight-years of GPS collar data were used to analyse the similarity in seasonal utilisation distributions for thirteen family groups. Elephants exhibited remarkable consistency in their seasonal range fidelity across the study with rainfall emerging as a key driver of space-use. Within years, high range fidelity from summer to autumn and from autumn to winter was driven by increased rainfall and the retention of high-quality vegetation. Across years, sequential autumn seasons demonstrated the lowest levels of range fidelity due to inter-annual variability in the wet to dry season transition, resulting in unpredictable resource availability. Understanding seasonal space use is important for determining the effects of future variability in environmental conditions on elephant populations, particularly when it comes to management interventions. Indeed, over the coming decades climate change is predicted to drive greater variability in rainfall and elevated temperatures in African savanna ecosystems. The impacts of climate change also present particular challenges for elephants living in fragmented or human-transformed habitats where the opportunity for seasonal range shifts are greatly constrained.

Skin wrinkles and folds enable asymmetric stretch in the elephant trunk.

The elephant's trunk is multifunctional: It must be flexible to wrap around vegetation, but tough to knock down trees and resist attack. How can one appendage satisfy both constraints? In this combined experimental and theoretical study, we challenged African elephants to reach far-away objects with only horizontal extensions of their trunk. Surprisingly, the trunk does not extend uniformly, but instead exhibits a dorsal "joint" that stretches 15% more than the corresponding ventral section. Using material testing with the skin of a deceased elephant, we show that the asymmetry is due in part to patterns of the skin. The dorsal skin is folded and 15% more pliable than the wrinkled ventral skin. Skin folds protect the dorsal section and stretch to facilitate downward wrapping, the most common gripping style when picking up items. The elephant's skin is also sufficiently stiff to influence its mechanics: At the joint, the skin requires 13 times more energy to stretch than the corresponding length of muscle. The use of wrinkles and folds to modulate stiffness may provide a valuable concept for both biology and soft robotics.

Cardiopulmonary responses of free-ranging African elephant (Loxodonta africana) bulls immobilized with a thiafentanil-azaperone combination.

OBJECTIVE: To determine the time course and certain cardiopulmonary effects of trunk-breathing elephants immobilized with thiafentanil-azaperone. STUDY DESIGN: Prospective descriptive study. ANIMALS: A convenience sample of 10 free-ranging African elephant bulls (estimated weight range: 3000-6000 kg). METHODS: Elephants were immobilized using thiafentanil (15-18 mg) and azaperone (75-90 mg) administered by dart. Once recumbent, the respiratory rate, minute ventilation (V˙e), end-tidal carbon dioxide (Pe'CO2), arterial blood pressure and heart rate were recorded immediately after instrumentation and at 5 minute intervals until 20 minutes. Arterial blood gases were analysed at the time of initial instrumentation and at 20 minutes. On completion of data collection, thiafentanil was antagonized using naltrexone (10 mg mg-1 thiafentanil; administered intravenously). A stopwatch was used to record time to recumbency (dart placement to recumbency) and time to recovery (administration of antagonist to standing). Data were compared using a one-way anova. Data are presented as mean ± standard deviation. RESULTS: All elephants were successfully immobilized, and there were no significant changes in cardiopulmonary variables over the monitoring period. Average time to recumbency was 12.5 (± 3.9) minutes. The measured V˙e was 103 (± 30) L minute-1. The average heart and respiratory rates over the 20 minute immobilization were steady at 49 (± 6) beats minute-1 and 5 (± 1) breaths minute-1, respectively. The mean arterial blood pressure was 153 (± 31) mmHg. The elephants were acidaemic (pH: 7.18 ± 0.06), mildly hypoxaemic (PaO2: 68 ± 15 mmHg; 9.1 ± 2.0 kPa) and hypercapnic (PaCO2: 52 ± 7 mmHg; 6.9 ± 0.9 kPa). Average time to recovery was 2.2 ± 0.5 minutes. CONCLUSION AND CLINICAL RELEVANCE: African elephant bulls can be successfully immobilized using thiafentanil-azaperone. Recumbency was rapid, the cardiopulmonary variables were stable over time, and recovery was rapid and complete. Mild hypoxaemia and hypercapnia were evident.

Developing welfare parameters for African elephants (Loxodonta africana) in fenced reserves in South Africa.

South Africa has many fenced reserves harbouring small to medium sized populations of African elephant (Loxodonta africana), most of which have been translocated. Elephants on fenced reserves may be exposed to various management interventions and practices (translocation, hunting, darting, high tourism impact, contraception programs, disruption due to infrastructure maintenance, etc.). These factors may impact the welfare of elephants. Poor elephant welfare may have serious consequences such as increased inter- and intra-species aggression that could result in fatalities. This is the first study to attempt to define behavioural and physiological welfare parameters for free-ranging elephants on small to medium sized reserves. The eight study sites incorporated reserves with different social structure combinations, elephant life-histories, reserve sizes, habitat, management, and tourism intensity. Data collection consisted of behavioural observations (10-minute videos) as well as faecal samples. By incorporating both behavioural and physiological (faecal glucocorticoid metabolite (fGCM) concentration) parameters, we aimed to investigate whether the two parameters showed similar trends. Five behavioural categories were identified (Arousal, Assessing, Ambivalent, Ambivalent/ Body care, and Frustrated behaviour), with various detailed behaviours demonstrated by the elephants that may indicate the influence of anthropogenic disturbance and possibly impact on animal welfare. The study showed significant differences between the selected detailed behaviours, behavioural categories and fGCM concentrations of elephants across the eight reserves. History seemed to be a decisive factor, as reserves with predominantly ex-captive elephants showed higher frequencies of certain behaviours as well as higher fGCM concentrations. Age, sex, reserve size and season were also found to contribute to our defined welfare indices and fGCM concentrations. This indicates that behavioural parameters, indicative of certain behavioural states, are valuable indicators of welfare, as supported by the physiological response of the elephants. The results also highlight the importance of taking multiple specified behaviours from a category into consideration when evaluating the welfare of elephants, to account for individual variation.

Effects of between and within Herd Moves on Elephant Endotheliotropic Herpesvirus (EEHV) Recrudescence and Shedding in Captive Asian Elephants (Elephas maximus).

Haemorrhagic disease associated with elephant endotheliotropic herpesvirus (Elephantid herpesvirus, EEHV) infections is the leading cause of death for Asian elephant (Elephas maximus) calves. This study assessed the effect of captive herd management on EEHV shedding, as evidence of latent infection reactivation, focusing on: (1) the influence of social change on the odds of recrudescence; (2) the respective effects of between and within herd moves; and (3) characteristics of recrudescent viral shedding. Trunk and conjunctival swabs (n = 165) were obtained from six elephants at an EAZA-accredited zoo, collected during a period of social stability, and at times of social change. Longitudinal sampling took place at times of moving two bulls out of the collection and one new bull into an adjacent enclosure to the cow herd (between herd moves), and during a period of mixing this new bull with the cow herd to facilitate mating (within herd moves). Quantitative PCR was employed to detect EEHV 1a/b, 4a/b, and EF-1-α (housekeeping gene). Generalised estimating equations determined EEHV recrudescence odds ratios (OR) and relative viral DNA load. Sixteen EEHV 1a/b shedding events occurred, but no EEHV 4a/b was detected. All management-derived social changes promoted recrudescence (social change OR = 3.27, 95% CI = 0.412-26, p = 0.262; and between herd moves OR = 1.6, 95% CI = 0.178-14.4, p = 0.675), though within herd movements posed the most significant increase of EEHV reactivation odds (OR = 6.86, 95% CI = 0.823-57.1, p = 0.075) and demonstrated the strongest relative influence (post hoc Tukey test p = 0.0425). Shedding onset and magnitude ranged from six to 54 days and from 3.59 to 11.09 ΔCts. Differing challenges are associated with between and within herd movements, which can promote recrudescence and should be considered an exposure risk to naïve elephants.

Simulated poaching affects global connectivity and efficiency in social networks of African savanna elephants-An exemplar of how human disturbance impacts group-living species.

Selective harvest, such as poaching, impacts group-living animals directly through mortality of individuals with desirable traits, and indirectly by altering the structure of their social networks. Understanding the relationship between disturbance-induced, structural network changes and group performance in wild animals remains an outstanding problem. To address this problem, we evaluated the immediate effect of disturbance on group sociality in African savanna elephants-an example, group-living species threatened by poaching. Drawing on static association data from ten free-ranging groups, we constructed one empirically based, population-wide network and 100 virtual networks; performed a series of experiments 'poaching' the oldest, socially central or random individuals; and quantified the immediate change in the theoretical indices of network connectivity and efficiency of social diffusion. Although the social networks never broke down, targeted elimination of the socially central conspecifics, regardless of age, decreased network connectivity and efficiency. These findings hint at the need to further study resilience by modeling network reorganization and interaction-mediated socioecological learning, empirical data permitting. The main contribution of our work is in quantifying connectivity together with global efficiency in multiple social networks that feature the sociodemographic diversity likely found in wild elephant populations. The basic design of our simulation makes it adaptable for hypothesis testing about the consequences of anthropogenic disturbance or lethal management on social interactions in a variety of group-living species with limited, real-world data.

Putative neural consequences of captivity for elephants and cetaceans.

The present review assesses the potential neural impact of impoverished, captive environments on large-brained mammals, with a focus on elephants and cetaceans. These species share several characteristics, including being large, wide-ranging, long-lived, cognitively sophisticated, highly social, and large-brained mammals. Although the impact of the captive environment on physical and behavioral health has been well-documented, relatively little attention has been paid to the brain itself. Here, we explore the potential neural consequences of living in captive environments, with a focus on three levels: (1) The effects of environmental impoverishment/enrichment on the brain, emphasizing the negative neural consequences of the captive/impoverished environment; (2) the neural consequences of stress on the brain, with an emphasis on corticolimbic structures; and (3) the neural underpinnings of stereotypies, often observed in captive animals, underscoring dysregulation of the basal ganglia and associated circuitry. To this end, we provide a substantive hypothesis about the negative impact of captivity on the brains of large mammals (e.g., cetaceans and elephants) and how these neural consequences are related to documented evidence for compromised physical and psychological well-being.

Anthropogenic pressures increase extinction risk of an isolated Asian elephant (Elephas maximus) population in southwestern China, as revealed by a combination of molecular- and landscape-scale approaches.

Identification of the effect of anthropogenic threats on ecosystem is crucial. We used molecular tools and remote sensing to evaluate the population status of an isolated Asian elephant population in southwestern China in response to changes in habitat suitability between 1989 and 2019. A total of 22 unique genotypes were identified from 117 dung samples collected between March and June 2018 using microsatellite DNA analysis, including 13 males and 9 females. Based on the size of fecal boli, 1 animal was a juvenile, 9 were subadults, and 12 were adults, indicating that recruitment was limited. The effective population size was small (15.3) but there was no signature of a recent population bottleneck. We observed a low genetic diversity (He = 0.46 ± 0.05) and a high level of inbreeding (Fis of 0.43 ± 0.11), suggesting low population viability and high risk of extinction. In total, these elephants lost nearly two thirds (62%) of their habitat in 3 decades. The expansion of agriculture and rubber plantations followed by an increase in human settlements after 1989 increased the isolation of this population. We recommend that resettlement of 800 inhabitants of 2 villages and the abandonment of associated farmland and rubber plantations would make an additional 20 km2 of suitable habitat available. This could allow a population increase of 14 elephants, possibly by translocating individuals from elsewhere in China. Our findings can be applied to the management and conservation of other fragmented populations in China or in other range countries of Asian elephants.

At high stocking rates, cattle do not functionally replace wild herbivores in shaping understory community composition.

Over a quarter of the world's land surface is grazed by cattle and other livestock, which are replacing wild herbivores, potentially impairing ecosystem structure, and functions. Previous research suggests that cattle at moderate stocking rates can functionally replace wild herbivores in shaping understory communities. However, it is uncertain whether this is also true under high stocking rates and the effects of wild herbivore on plant communities are moderate, enhanced, or simply additive to the effects of cattle at high stocking rates. To evaluate the influence of cattle stocking rates on the ability of cattle to functionally replace wild herbivores and test for interactive effects between cattle and wild herbivores in shaping understory vegetation, we assessed herbaceous vegetation in a long-term exclosure experiment in a semi-arid savanna in central Kenya that selectively excludes wild mesoherbivores (50-1000 kg) and megaherbivores (elephant and giraffe). We tested the effects of cattle stocking rate (zero/moderate/high) on herbaceous vegetation (diversity, composition, leafiness). We also tested how those effects depend on the presence of wild mesoherbivores and megaherbivores. We found that herbaceous community composition (primary ordination axis) was better explained by the presence/absence of herbivore types than by total herbivory, suggesting that herbivore identity is a more important determinant of community composition than total herbivory at high cattle stocking rates. The combination of wild mesoherbivores and cattle stocked at high rates led to increased bare ground and annual grass cover, reduced perennial grass cover and understory leafiness, and enhanced understory diversity. These shifts were weaker or absent when cattle were stocked at high stocking rates in the absence of wild mesoherbivores. Megaherbivores tempered the effects of cattle stocked at high rates on herbaceous community composition but amplified the effects of high cattle stocking rate on bare ground and understory diversity. Our results show that cattle at high stocking rates do not functionally replace wild herbivores in shaping savanna herbaceous communities contrary to previous findings at moderate stocking rates. In mixed-use rangelands, interactions between cattle stocking rate and wild herbivore presence can lead to non-additive vegetation responses with important implications for both wildlife conservation and livestock production.